-
THE ALFRED LEEDS FOSSIL VERTEBRATE COLLECTION OF THE NATIONAL
MUSEUM OF IRELAND—NATURAL HISTORY
RICARDO ARAÚJO, ADAM S. SMITH1 AnD JEFF LISTOn
(Received 05 October 2007. Accepted 14 August 2008.)
Abstract
Alfred nicholson Leeds (1847–1917) is famous among vertebrate
palaeontologists for amassing an invaluable collection of fossil
vertebrates from the Middle Jurassic aged ‘Oxford Clay’ deposits of
the Peterborough district in the UK, throughout the late nineteenth
and early twentieth centuries. Part of his collection was acquired
by the national Museum of Ireland—natural History in november 1893
but has not previously been described. This fossil material
includes a suite of Jurassic marine reptiles: crocodiles,
plesiosaurs and ichthyosaurs. There are no examples, however, of
the giant fish Leedsichthys; the remains of which are commonly
found among the fauna of the Oxford Clay Formation. Despite
representing only a very small fraction of the total Leeds
Collection, the specimens in Ireland are significant historically,
and have scientific value in their own right. For the first time,
in this paper, a historical review of the Dublin Leeds Collection
is presented, the fossil specimens described and a long-lost piece
of important documentation is presented.
Historical review of the Leeds Collection
Since the nineteenth century the clay pits of the Peterborough
district, UK, have yielded a huge number of marine reptile remains.
The collection housed in the nMInH (national Museum of
Ireland—natural History), although a very small fraction of the
Leeds Collection in total, is valuable because of its historical
relevance and the intrinsic scientific value of its fossils.
Although primarily known for its Callovian age marine reptile
skeletons, the Oxford Clay also yields fish, including the giant
suspension-feeder Leedsichthys (Liston and noè 2004), and dinosaurs
(Liston and noè 2008). Alfred Leeds also included coprolites and
invertebrate fossils (e.g. bivalves like Trigonia, belemnites and
ammonites) in many of his sales, to
provide institutions with a representative faunal suite for the
Oxford Clay. General overviews of the Leeds Collection were
undertaken by Andrews (1910) and Leeds (1956).
The collection was amassed by Alfred Leeds between c. 1867 and
1917 (Leeds 1956) when he assumed the management of his family’s
farm in the Peterborough district, Cambridgeshire. Charles Leeds,
his elder brother, occupied a minor role in the collection of
specimens before he emigrated to new Zealand in 1887. Charles was
originally encouraged by Professor John Phillips, Lecturer in
Geology at Oxford University, to preserve and continue collecting
fossils from the Peterborough brick pits (Woodward 1917) and
Phillips was the first palaeontologist to publish a paper on
specimens in the collection
1Research related to this paper was carried out while the author
was affiliated to the School of Biology and Environmental Science,
University College Dublin.
Irish Journal of Earth Sciences 26 (2008), 17–32 © Royal Irish
Academy
17
Araujo.indd 17 14/01/2009 15:21:18
-
18 Irish Journal of Earth Sciences (2008)
(Phillips 1871), describing plesiosaur and pliosaur remains.
Another isolated encounter between the collection and professional
palaeontologists came in 1874, when the well-known fossil reptile
specialist Harold (Harry) Govier Seeley came to Eyebury—the Leeds’
residence and fossil store (Leeds 1956)—and subsequently published
two papers (Seeley 1874a, 1874b) on partial skeletons of new genera
in the collection. But it was not until almost twenty years after
the inauguration of the Leeds Fossil Collection that it came to the
attention of the general scientific community, when in 1885 Alfred
Leeds first contacted Henry Woodward, Keeper of Geology in the
British Museum (natural History),– BMnH. The surprising lack of
further visits during this early period has been commented on at
length by Leeds’ son Edward Thurlow Leeds in his history of the
collection (Leeds 1956)—it was clear that this was a new and
significant resource, yet no one appears to have responded to the
publications by Phillips and Seeley by seeking out this collection
for themselves. As E.T. Leeds comments, it may be down to the lack
of inclination of the Leeds brothers to approach anyone, perhaps
because in the early days of the collection they could obtain the
information that they needed from publicly available books.
Following Alfred Leeds’ contact, Henry Woodward visited the
collection at Eyebury, an event that was to prove noteworthy for
two reasons: (1) his visit spurred other palaeontologists to follow
(R. Lydekker, J. W. Hulke and even O. C. Marsh); and (2) the
friendly relationship which developed between H. Woodward and A.
Leeds became fruitful for the BMnH, the institution now boasting
the choicest parts of the collection (Woodward 1917). Woodward
became interested in acquiring this collection (cited in Leeds
1956) and by 1890, five years after his initial visit, the BMnH
agreed to purchase the items (about 5,000kg of bones). The Leeds
Collection is nowadays dispersed throughout Britain (e.g. the
Hunterian Museum, University of Glasgow, houses the bulk of the
collection, over 600 specimens purchased from Leeds’ son, Edward
Thurlow, and the Royal Scottish Museum in Edinburgh has some
material obtained via Stürtz (see below)); Germany (e.g. Tübingen
and the Goldfuss Museum of Bonn); Austria; France; Sweden
(Uppsala); the USA (Peabody Museum, Yale University) and Ireland
(Woodward 1917; neaverson 1935; Liston and noè 2008; this paper).
The purpose here is to outline the history of the Dublin Leeds
Collection, and provide a catalogue of the specimens together with
additional anatomical information and illustrations where
appropriate.
The Dublin Leeds Collection
The correspondence files of the nMInH show how the relationship
between Alfred Leeds and the Dublin Science and Art Museum (as the
nMInH was then known) developed. As recorded by Alfred Leeds, he
first made an approach to the institution in October 1892, asking
if it would like to ‘purchase some fossil bones’ (Alfred Leeds to
Robert Scharff, 3 August 1893—nMInH SA Letter File, 1893, Vol.3;
note that all references in this section to Leeds correspondence
refer to A.n. Leeds, unless otherwise stated. All correspondence
referenced in this article is summarised in Table 1). At the time,
the Science and Art Museum informed him that their annual purchase
grant had
Table 1—List of correspondence referenced within this paper,
ordered by appearance.
Archival reference Details Date
nMInH SA Letter File, 1893, Vol. 3 Alfred Leeds to Valentine
Ball 22 October 1893
nMInH SA Letter File, 1893, Vol. 3, no. 508
1893
nMInH SA Letter File, 1893, Vol. 3 Alfred Leeds to Valentine
Ball 12 October 1893
nMInH Letter File, 1893, Vol. 3 Valentine Ball to Alfred Leeds
21 October 1893
nMInH Minute Paper by Robert Francis Scharff, Keeper of natural
History at nMInH
9 november 1893
nMInH 1896.L Correspondence Files 1896
nMInH SA Letter File, 1893, Vol. 3 Alfred Leeds to Robert Sharff
3 August 1893
nMInH SA Letter File, 1893, Vol. 3 Alfred Leeds (informal list
of the specimens) 1893
Araujo.indd 18 14/01/2009 15:21:19
-
Araújo et al.—The Alfred Leeds Collection of the NMINH 19
been spent, but that he should renew his offer in the future.
Alfred Leeds repeated his offer in August 1893, inviting museum
representatives to visit his collection to view what he was
offering, and he pointed out how conveniently located he was with
respect to journeys from either London which was one and a half
hours away or nottingham, which took one-quarter of an hour less
(Alfred Leeds to Robert Scharff, 3 August 1893—nMInH SA Letter
File, 1893, Vol.3).
Within two days, a discussion ensued between Dr Robert Francis
Scharff, Keeper of natural History, and Dr Valentine Ball, Director
of the then Dublin Science and Art Museum (now nMInH), both
enthusiastic to procure items, and in agreement that Ball should
visit Eyebury from nottingham (Minute Paper between Valentine Ball
and Robert Scharff, 5–7 november 1893). Implicit within this
dialogue is that Ball had already arranged to attend a meeting in
nottingham prior to Alfred Leeds’ letter, and although it is very
likely that it was the Sixty-Third Meeting of the British
Association for the Advancement of Science, held at nottingham in
September 1893 (British Association for the Advancement of Science
1894), no information is given to corroborate this. A telegram was
sent by Ball to Scharff from the Conference Hall in nottingham,
arriving in Dublin on the 21 September 1893, announcing that he was
‘going to see the bones’ and asking if it might be possible to
spend up to £50 if it was ‘desirable’ to do so—presumably this was
dependent on his imminent assessment of the material at Eyebury
(nMInH SA Letter File, 1893, Vol. 3). Ball’s visit must indeed have
been brief, as he does not sign the Eyebury Visitors’ Book for the
end of September (when Henry Woodward, Keeper of Geology in the
BMnH and his wife were visiting); but the material must indeed have
been ‘desirable’, as a request for funding from Ball reached the
Lords of the Committee of Council on Education at the Department of
Science and Art in London the next day, and by the 26 September
they had given their sanction to spending £70 on ‘Bones of saurians
&c.’ for the Dublin Museum of Science and Art (26 September
1893, nMInH SA Letter File, 1893, Vol. 3, no. 508).
As in the previous year, however, budgetary constraints
intervened, and by the 5 October 1893 Ball informed Alfred Leeds
that £35 could be paid, with the balance forthcoming after 1 April,
presumably reflecting the commencement of the institution’s next
financial year. Alfred Leeds appeared undeterred by this payment in
instalments, and by 22 October dispatched a ‘holding’ letter to
Ball, informing him that Leeds had been away from home and thus was
delayed in completing the packing of the material (Alfred Leeds to
Valentine Ball,
12 October 1893—nMInH SA Letter File, 1893, Vol. 3). On the 2
november Leeds sent Ball a further letter to confirm that all the
specimens were packed with details of how to extricate a mounted
ichthyosaur paddle that he had secured within one crate, and
expressing the hope that everything would ‘travel safely’ (Alfred
Leeds to Valentine Ball, 2 november 1893—nMInH SA Letter File,
1893, Vol. 3). He also indicated that he had forgotten to include
belemnites, but would send them as soon as he could. The material
was entered in the accession register by Scharff on 9 november 1893
(nMInH 1893.156–74), and he certified the accounts the same day as,
although an Asteracanthus dorsal spine had not been sent, it had
been intended as a gift rather than a purchased item (Minute Paper
by Robert Francis Scharff to Valentine Ball, 9/11/1893). Alfred
Leeds wrote again to Ball to express his relief at the safe arrival
of the consignment, as he,
was very much afraid that the heads might come to grief on the
journey—I think that I could not have put the spine of
Asteracanthus in the box. I know I picked one out to send—but
certainly did not put it in the little box—so one shall be looked
out and sent you by post. I sent some belemnites by post last week
& hope you have received them.
(Alfred Leeds to Valentine Ball, 12 november 1893—nMInH SA
Letter File, 1893, Vol. 3)
Although in his letter of 2 november 1893 Alfred Leeds indicated
that if his list of specimens did ‘not tell you all you want to
know about the specimens—I shall be glad to answer any questions I
can—I think I have put in all the bones agreed upon by us—if not
please let me know’ (Alfred Leeds to Valentine Ball, 2 november
1893—nMInH SA Letter File, 1893, Vol. 3), it seems that the staff
at Dublin were slow to take him up on his offer. This may have been
because a recent policy of employing external curators to catalogue
the fossil collections had lapsed, and there was simply not the
staff to keep on top of new material as it came in. Whatever the
reason, it was three years before an enquiry arrived asking for
more information, including whether or not the specimens had been
described, and Alfred Leeds replied (Letter from Alfred Leeds, 28
October 1896—nMInH, October 1896 Correspondence File) indicating
that he could not remember what he had sent, and had kept no
record. He offered some crocodile remains for sale (‘I believe I
get nine or ten distinct species’) but no further purchases appear
to have been made by the Dublin Museum of Science and Art (Letter
from Alfred Leeds, 28th October 1896—nMInH, October 1896
Correspondence File).
Araujo.indd 19 14/01/2009 15:21:20
-
20 Irish Journal of Earth Sciences (2008)
The Dublin Leeds Collection is noteworthy because it was the
first suite of material sold by Alfred Leeds to an institution
other than the BMnH, prior to Leeds establishing a relationship
with the Bonn dealer Bernhard Stürtz in 1897. One clear drawback of
this collection, however, is that no precise information about the
geological horizon was provided for any of the Dublin Leeds
specimens. Andrews (1910) states that ‘the horizon at which [Oxford
Clay] reptilian bones occur is that characterised by the presence
of the “Ornatus” group of Ammonites’ (p. vi), particularly
Cosmoceras gulielmii. Arkell (1933 in Duff 1975) further specified
the zone in which reptile remains are found, to Kosmoceras jason
(Middle Callovian). The Dublin Leeds Collection therefore probably
originated from this zone of the ‘Lower Oxford Clay’. Although
precise stratigraphic positions can retrospectively be determined
for some of Alfred Leeds’ specimens from his excavation records in
the ‘Eyebury Register’, the only notes relating to the Dublin
specimens do not contain salient information.
Systematic palaeontology
Crocodylomorpha Hay, 1930 (sensu Walker, 1970)Crocodyliformes
Hay, 1930 (sensu Benton and Clark, 1988)Thalattosuchia Fraas,
1901Metriorhynchidae Fitzinger, 1843Metriorhynchus Meyer, 1830
Metriorhynchus superciliosus (de Blainville, 1853)
Eudes-Deslongchamps, 1867–9
MaterialSpecimen nMInG F16892 (Fig. 1A) comprises a skull
(almost complete) and lower jaw, associated with 16 vertebrae.
Specimen nMInG F21731 (Fig. 1B) consists of the skull (almost
complete) and the lower jaw, 28 vertebrae and both ilia. The skull
has been slightly dorso-ventrally compressed. The dorsal skull
bones bear smooth ornamentation and dorsal crests occur on the
parietal and squamosal. nMInG F21760 is an isolated left femur.
The skulls of the nMInG F16892 and nMInG F21731 measure 600mm
and 670mm respectively, according to Vignaud’s (1995) standardised
measurements
Fig. 1—Skull of Metriorhynchus (in dorsal view) and both lower
jaw rami (in lateral view). A. Specimen nMInG F16892; B. Specimen
nMInG F21731. Scale-bar equals 30cm.
Araujo.indd 20 14/01/2009 15:21:40
-
Araújo et al.—The Alfred Leeds Collection of the NMINH 21
and the nasal bones are 230mm (nMInG F16892) and 255mm (nMInG
F21731) long. The supratemporal fossa is 115mm (nMInG F16892) and
140mm (nMInG F21731) long, 60mm and 90mm wide respectively. The
lower jaw of the nMInG F21731 measures 715mm in length, the
mandibular symphysis is 400mm long and the mandibular tooth row
spans 335mm.
DiscussionBoth the Metriorhynchus skulls were prepared in 2004
at University College Dublin; F16892 was originally embedded in
paraffin wax and only the dorsal surface of the skull was exposed
(Gandola et al. 2006). The associated postcranial material is
generally well-preserved despite some vertebrae lacking neural
arches. The skull of nMInG F21731 is broken into five pieces, since
it was originally repaired with natural animal glue. The specimen
exhibits dorso-ventral flattening post mortem deformation. Specimen
nMInG F21731 has dorsal crests on the parietal and squamosal, and
pitted ornamentation on the frontal and prefrontals. These
characters combined with the skull proportions are diagnostic for
M. superciliosus (Vignaud 1995; Adams-Tresman 1987a). The
postcranial material—28 vertebrae—are broken into pieces. The
taphonomy of this specimen is interesting since anterior and
posterior remains of the specimen are found together. An
interesting anatomical feature in Metriorhynchus, salt glands in
the antorbital cavity, was first detected in these specimens
(Gandola et al. 2006).
The isolated left femur nMInG F21760 was initially suggested by
A. Leeds as also belonging to Metriorhynchus (informal list of the
specimens by Alfred Leeds—nMInH SA Letter File, 1893, Vol. 3). This
femur is similar to Steneosaurus, but one autapomorphic character,
an extended ventrally roughened trochanter, is absent in
Steneosaurus.
Teleosauridae Geoffroy, 1831Steneosaurus Geoffroy, 1825
Steneosaurus edwardsi (Eudes-Deslongschamps, 1868)
Eudes-Deslongchamps, 1867–9 sensu Vignaud, 1995
Material Specimen nMInG F21732 (Fig. 2A) comprises the anterior
part of the skull (Fig. 3A) and mandible (~60% of total), two
isolated teeth, tarsal bones (a right calcaneum and a right fifth
metatarsal), 16 vertebrae including the atlas-axis (Fig. 3B) and
rib fragments. Specimens nMInG F21762, nMInG F21763 are deduced to
be additional teeth from two different individuals, due to the
different preservation.
The measurements are standardised from Vignaud (1995). The total
length of the skull is estimated as 900mm, with 270mm between the
premaxillary bone and the anterior end of the nasal. The maximum
length of the orbit is 60mm and its perpendicular length is 45mm.
The width of the skull measured on the anterior edge of the orbits
is estimated as 150mm, while the atlas-axis complex (including the
atlantal ring) measures 73mm in length, 40mm in width and 9mm in
height (estimated). The preserved cervical vertebrae range in
central length from 46mm to 49mm and in width from 36mm to 47mm.
The dorsal centra range from 50mm to 56mm in length, 44mm to 48mm
in width, and including the neural arch (from the vertebrae where
preserved) are 95mm to 103mm in height.
DiscussionThe most striking feature of nMInG F21732 is its long
narrow snout (Fig. 3A). In addition, the square-like supratemporal
fenestrae, the large forward pointing external nares, the long
mandibular symphysis and the latero-dorsally orientated
sub-circular orbits are important autapomorphies of the genus
Steneosaurus (Adams-Tresman 1987b; Vignaud 1995). We have attempted
to assign nMInG F21732 on the basis of our anatomical observations.
Some species can be readily excluded—S. bollensis, S. leedsi and S.
megarhinus—because they share the longirostrine condition and have
finely built skulls (Andrews 1913). Among the most well-preserved
brevirostrine taxa, S. obtusidens, S. edwardsi and S. bouchardi are
the best candidates for this assignment. S. obtusidens, however,
possesses a different dental formula—although this is a very weak
character since it can vary intraspecificially—in the lower jaw and
the characteristic pointed and curved shape of the teeth of S.
obtusidens is not detected in nMInG F21732. Steneosaurus bouchardi
has never been reported from the Jurassic of the UK, and, unlike
nMInG F21732 belongs to the Upper Kimmeridgian. This crocodile also
has a heavily built skull; its supratemporal fossae are rectangular
and it has a marked antorbital foramen (Vignaud 1995). On the basis
of these arguments, S. edwardsi seems to be the most plausible
species for referral of nMInG F21732; not only are the proportions
of the skull similar, but also the dental formula, the absence of
the antorbital foramen and the circular shape of the orbits are
characteristics shared by nMInG F21732. The S. edwardsi specimen
BMnH R2075—collected from the same general locality—stored in the
natural History Museum (London) also exhibits a very similar
preservation to nMInG F21732.
Araujo.indd 21 14/01/2009 15:21:42
-
22 Irish Journal of Earth Sciences (2008)
Fig. 2—Outline of three marine reptiles’ skeletons, showing the
elements preserved in the most complete specimens (shaded grey)
within the Dublin Leeds Collection. A. Steneosaurus edwardsi, in
dorsal view (nMInG F21732); B. Cryptoclidus eurymerus, in lateral
view (nMInG F21786); C. Ophthalmosaurus icenicus, in lateral view
(nMInG F21787). A is based on a Teleosaurus from Steel (1973). B
and C are based on Andrews (1910). not to scale.
Araujo.indd 22 14/01/2009 15:21:53
-
Araújo et al.—The Alfred Leeds Collection of the NMINH 23
Fig. 3—Specimen nMInG F21732 ascribed here to Steneosaurus
edwardsi, some bones preserving important diagnostic features. A.
Skull in dorsal view; B. Atlas-axis complex in lateral view.
Scale-bar in A equals 20cm and in B equals 1cm.
Araujo.indd 23 14/01/2009 15:22:05
-
24 Irish Journal of Earth Sciences (2008)
The vertebrae of nMInG F21732 are all slightly distorted, so
that in six of them the anterior articular facet is sheared to the
left, relative to the posterior articular facet, and in seven the
anterior facet is displaced to the right. This indicates that the
neural spine was not in close articulation in situ, and may also
explain why so much of the rest of the skeleton is missing (i.e.
due to dispersal on the seabed prior to burial). The element
identified as a fifth metatarsal is uncertain due to the poor
preservation of the bone.
Plesiosauria de Blainville, 1835Plesiosauroidea Gray,
1825Cryptoclididae Williston, 1925Cryptoclidus Seeley, 1892
Cryptoclidus eurymerus (Phillips 1871) Seeley, 1892
MaterialThe remains of four specimens of Cryptoclidus are
included in the Dublin Leeds Collection. The most complete
individual is nMInG F21786 (Fig. 2B) which comprises a large
proportion of the vertebral column with neural arches, both humeri
(each 260mm long, with a distal width of 155mm), radii (85mm long),
ulnae (70mm long), a number of carpals and metacarpals, a complete
right scapula (190mm long) and the dorsal process of the left
scapula, the anterior part of the left coracoid (preserved part
175mm long), a partial clavicle (not figured), a partial pubis
(205mm long) and associated rib fragments (Fig. 4).
Specimen nMInG F21785 belongs to a larger individual and
comprises the distal part of a humerus (305mm wide distally), a
radius, two ulnae and a partial ischium (maximum width 230mm), two
phalanges,
eleven vertebrae, and associated rib fragments. The preservation
of this specimen contrasts sharply with most of the other material
in the Dublin Leeds Collection; it is likely that this specimen
originates from a distinct locality and/or horizon.
Specimen nMInG F21782 is an isolated right humerus, partly
restored in plaster and almost identical in size and form to the
right humerus of nMInG F21786.
Specimen nMInG F21781 is the poorly developed humerus of a
juvenile plesiosaur, probably also referable to Cryptoclidus due to
its general morphology.
DiscussionCryptoclidus is widely considered to be the most
abundant plesiosaur in the Oxford Clay Formation (Martill 1991) and
this is certainly the case within the small Dublin collection. The
Cryptoclidus material in the Dublin Leeds Collection ranges from
juvenile to old adult individual (sensu Brown 1981).
Specimen nMInG F21785 is a large specimen of Cryptoclidus in
which the preserved parts of the forelimb (humerus, radius, ulna)
are diagnostic (Brown 1981)—the radius and ulna have a distinctive
shape and there is a clear ‘elbow’ formed by the angle between the
anterior border of the humerus and radius. A small bone fused to
the postaxial margin of the ulna is of particular interest (this
bone is separate in the second limb but the tight suture shows that
it has been broken off post-fossilisation); although labelled as an
ulnare, its position is inconsistent with this diagnosis and
following Caldwell (1997) should rather be regarded as the
pisiform. Fusion of the pisiform occurs late in the ontogeny of
this genus and is a character seen in
Fig. 4—The partial pectoral girdle and forelimbs of Cryptoclidus
eurymerus (nMInG F21786) in dorsal view. The full width of the
girdle has been reconstructed based on the complete left coracoid.
The organisation of the bones in the limbs was based on Caldwell
(1997). Scale-bar equals 30cm.
Araujo.indd 24 14/01/2009 15:22:09
-
Araújo et al.—The Alfred Leeds Collection of the NMINH 25
‘Apractocleidus’ (Smellie 1916), a genus regarded by Brown
(1981) as an ‘old adult’ Cryptoclidus. The fused pisiform, fused
neural arches and general large size corroborate the status of
nMInG F21785 as an old individual.
On the other hand, specimen nMInG F21786 shows many juvenile
characters (Andrews 1895)—a poorly developed pectoral girdle (the
anterior process of the scapula does not contact the midline),
poorly ossified limbs, and neural arches which are not ossified to
the centrum (Fig. 4). The radius of Cryptoclidus, however, is
diagnostic even in juveniles, ‘being greatly expanded proximally
and anteriorly’ (Brown 1981, 276).
Muraenosaurus Seeley, 1874a
Muraenosaurus sp.
MaterialSpecimen nMInG F21778 is an incomplete right scapula,
250mm long. The dorsal process is 140mm long on the lateral surface
and the ventral process is 150mm wide.
DiscussionThe ventral ramus of this incomplete right scapula
(nMInG F21778) is very broad and well-developed, as in old-adult
cryptoclidids, and appears to have met on the midline posteriorly.
Anteriorly, the ramus is excavated and separated from its
counterpart on the midline, a characteristic of Muraenosaurus
(Brown 1981). Apart from size, however, species of Muraenosaurus
cannot be differentiated based on scapular morphology alone (Brown
1981).
Muraenosaurus leedsii Seeley, 1874a
MaterialSpecimen nMInG F21779 is a single right femur (Fig. 5),
335mm long and 175mm wide distally.
DiscussionThere are two valid species of Muraenosaurus described
from the UK—M. leedsii and M. beloclis (Brown 1981). Specimen nMInG
F21779 is labelled as ‘M. plicatus’, a species now considered
synonymous with M. leedsii. Indeed, specimen nMInG F21779 most
closely approximates M. leedsii in the elongate and light
construction of the femur (Andrews 1910).
Pliosauroidea Seeley, 1874cPliosauridae Seeley, 1874cPeloneustes
Lydekker, 1889
Peloneustes philarchus (Seeley, 1869) Lydekker, 1889
MaterialTwo isolated propodial bones, nMInG F21783 (Fig. 6A) and
nMInG F21784 (Fig. 6B) and a tooth (nMInG F21761) belong to the
pliosaur Peloneustes philarchus.
DiscussionAccording to Tarlo (1960) the pliosaur Peloneustes is
monospecific. Its propodials are quite distinct from other
plesiosaur species in their bi-symmetrical tear-drop shaped outline
(see Fig. 6). The humerus of Peloneustes is much broader than the
femur (Andrews 1913) and on
Fig. 5—Right femur of Muraenosaurus beloclis (nMInG F21779).
Scale-bar equals 10cm.
Araujo.indd 25 14/01/2009 15:22:16
-
26 Irish Journal of Earth Sciences (2008)
this basis specimen nMInG F21784 may be a humerus, and nMInG
F21783 a femur. The tuberosity of nMInG F21784 is small, however,
and contradicts the usually strongly developed condition seen in
Peloneustes (Andrews 1913), perhaps indicating the immaturity of
the individual. The pattern of ornamentation on the isolated tooth
(nMInG F21761) is diagnostic (Martill 1991), therefore we assign
this specimen to Peloneustes.
Plesiosauria indet.
Material Specimen nMInG F21780 is a single complete ilium, 205mm
long.
DiscussionThe most notable feature of this isolated ilium is a
sharp kink in its shaft, which differs from the lightly curved ilia
seen in all other Oxford Clay Formation
plesiosaurs (see figures in Andrews 1910). Thus, this bone is
temporarily regarded as Plesiosauria indet. despite its apparently
distinctive morphology.
Ichthyopterygia Owen, 1840Ichthyosauria de Blainville,
1835Ophthalmosauridae Baur, 1887Ophthalmosaurus Seeley, 1874b
Ophthalmosaurus icenicus Seeley, 1874b
MaterialThis specimen (nMInG F21787) consists of an almost
complete pectoral girdle (Fig. 7), both humeri, 80 paddle bones
including one complete paddle (Fig. 8A), two cervical ribs, and a
femur (Fig. 2C). The scapulae are 220mm long and 125mm wide; the
minimum distance between the scapulae as articulated is 280mm. Each
humerus is 155mm long, and 140mm wide distally.
Fig. 6—Pelonestes philarchus. A. Femur (nMInG F21783); B.
Humerus (nMInG F21784). Scale-bars equal 10cm.
Araujo.indd 26 14/01/2009 15:22:26
-
Araújo et al.—The Alfred Leeds Collection of the NMINH 27
Fig. 7—Reconstructed pectoral girdle of Ophthalmosaurus icenicus
(nMInG F21787) in dorsal view. Scale-bar equals 20cm.
Fig. 8—Left paddle of Ophthalmosaurus icenicus (nMInG F21787) in
dorsal view. A. Reconstruction of the limb; B. Original in situ
illustra-tion by Alfred Leeds. Scale-bar equals 20cm.
Araujo.indd 27 14/01/2009 15:22:37
-
28 Irish Journal of Earth Sciences (2008)
DiscussionThe genus Ophthalmosaurus is characterised by a flared
proximal end of its scapula (Maisch and Matzke 2000) and in the
morphology of the humerus and phalanges, (McGowan and Motani 2003;
Sander 2000), all of which are preserved in nMInG F21787. According
to correspondence from A. Leeds (Letter from Alfred Leeds, 28
October 1896, Correspondence File 28/2/1896), the complete
Ophthalmosaurus paddle in the collection represents the ‘first
paddle of this species that was ever known to have all the bones in
the right position. I got each bone out and numbered it—& made
a drawing of the paddle as it lay in the clay’. This drawing was
not sent to Dublin, and during the course of this current work, the
illustration in question was traced (see Fig. 8). It was found to
still be in the possession of the Leeds family, as a part of Plate
23 of E.T. Leeds unpublished 1938/9 manuscript ‘Eyebury and the
Leeds Collection’, from which W.E. Swinton, as editor, produced the
posthumous volume The Leeds Collection of fossil reptiles in 1956.
The illustration is reproduced for the first time here (Fig. 8B),
and has proved invaluable in reconstructing a limb that would
otherwise be problematic as Ophthalmosaurus phalanges are neither
tightly packed nor close-fitting (McGowan and Motani 2003).
General discussion
The Museum of Science and Art in Dublin was established in 1877
when Ireland first took on responsibility for the funding and
management of Dublin museums. The existing museum of the Royal
Dublin Society was the primary contributor of palaeontological
collections to the new state museum (Monaghan 1992). This new
museum, however, was under the same administration as museums in
London and this had two effects. For existing collections there was
an increased professionalism that saw a number of external
palaeontologists employed by the Dublin Museum of Science and Art
as temporary curators to catalogue the existing fossil collections
(Monaghan 1992), namely: Davis (1888), Kidston (1888; Liston and
Sanders 2005) and Lydekker (1884, 1891). Lydekker’s catalogue of
the fossil vertebrate collection (1891) was in a similar style to
his work on the collections in the BMnH (e.g. Lydekker 1888–90).
This administrative arrangement also led to a funding regime that
allowed for significant purchases of new specimens. Valentine Ball,
Director of the state institutions of Science and Art in Dublin
from 1883–95, —and R.F. Scharff, — Keeper of natural History (as
well as other areas) from 1891–1921, were the personnel of the
nMInH involved
with the negotiation for and acquisition of the Leeds
Collection. The intention of acquiring large vertebrate fossils
seems to be connected to the palaeontological furore at the time,
but also complimented and built upon previous acquisitions of
fossil vertebrates (the William Lee Collection of Lias marine
reptiles for example). Richard Lydekker had identified gaps in the
fossil collections and had the contacts to assist in identifying
potential acquisitions.
There are four main issues related to the Dublin Leeds
Collection: (1) the missing material, (2) the numeration of the
specimens, (3) the two different shipments and their registration
process and (4) Ophthalmosaurus and the field illustration of its
paddle in situ.
Some parts of the collection are missing or in confusion.
namely, the above referred Asteracanthus dorsal spine, which was
not originally packed by Leeds in the shipment, but was registered
(nMInH 1893.179) on 20 november 1893, so presumably was sent
subsequently, as Alfred Leeds undertook to do in his letter of 12
november 1893 (Alfred Leeds to Valentine Ball, 12 november
1893—nMInH SA Letter File, 1893, Vol. 3). In addition, three teeth
are missing from a batch of five isolated teeth (accession number:
nMInH 1893.173). The two existing isolated teeth belong to
Peloneustes and Steneosaurus (see above). A plesiosaur ventral rib
(accession number: nMInH 1893.164) listed in the collection was
also not found in the nMInH storage. It may be that some specimens
that were painted with varnish (as a pyrite treatment) in 1979
could have subsequently been lost through pyrite decay. Four of the
phalangeal elements from Ophthalmosaurus Paddle 77 (nMInG F21787)
are missing, whether this occurred prior to packing, in transit, or
later during collection movement over the decades in Dublin is
unknown.
The numbering of the specimens can be confusing. Each has been
allocated three numbers—‘Leeds numbers’ were given by Leeds while
acquiring the fossils (marked with circular or oval labels). These
can be confirmed as such, as some are duplicated in the Eyebury
Register of the ‘Second’ Leeds Collection, where entries of
‘Dublin’ and ‘1892’ can be found (e.g. under the crocodiles heading
for entries ‘9’ and ‘12’). An ‘accession number’ was provisionally
applied to each specimen upon its arrival at the museum, and marked
with black ink directly on the bones (= the ‘registration number’
in the accession book). Some of the numbers are marked with white
ink: these were written on the bones in 1979 when they were crated,
for movement to a different storage facility. The actual official
registration numbers were only applied in 2007 for the purposes of
this paper (see Table 2).
Araujo.indd 28 14/01/2009 15:22:38
-
Araújo et al.—The Alfred Leeds Collection of the NMINH 29
Although the payment of £70 to Leeds was made in two instalments
of £35, this seems to have simply been a reflection of the
necessity of spreading the purchase over two financial years, as
all the material arrived in one shipment and was registered at the
same time (9 november 1893).
The discovery of the long-lost diagram of the paddle of
‘Ophthalmosaurus 77’ was a particularly satisfactory
outcome of this work. It seems most likely that the diagram was
never intended to be sent, either because of its slightly imperfect
nature (Alfred appears to have reworked four or five of the
outlines), or because he always intended to retain it as a guide
for himself to reconstruct further paddles of Ophthalmosaurus. It
seems most likely that Alfred Leeds produced the diagram as a
reference document to use when performing future
Table 2—List of the fossil marine reptiles in the Dublin Leeds
Collection, outlining the register numbers, taxo-nomic
identification and preserved elements.
Register number Taxonomic identification Preserved
element(s)
nMInG F21778 Muraenosaurus sp. Right scapula
nMInG F21779 Muraenosaurus leedsi Left femur
nMInG F21780 Plesiosauria indet. Right ilium
nMInG F21781 Cryptoclidus sp. Humerus (juvenile)
nMInG F21782 Cryptoclidus sp. Right humerus (juvenile)
nMInG F21783 Pelonestes philarchus Right femur
nMInG F21784 Pelonestes sp. Left humerus
nMInG F21759 Metriorhynchus sp. Left femur
nMInG Plesiosaurus Ventral rib—not located
nMInG F21760 Steneosaurus Scute
nMInG F21785 ‘Cimoliosaurus eurymerus’ (now Cryptoclidus
eurymerus)
1 left humerus (distal part), 1 left radius, 2 ulnae, 1 right
ischium, 2 phalanges, 11 vertebrae centra (1 complete), 3 rib
fragments
nMInG F21786 Cryptoclidus eurymerus 2 humeri, 2 radii, 2 ulnae,
25 vertebrae, 1 scapula, (left) half coracoid, 1 clavicle, half
pubis, 28 ribs, 24 neural arches
nMInG F21787 Ophthalmosaurus icenicus 2 humeri, 80 paddle bones,
2 coracoids, 2 scapulae, 3 clavicle fragments, 2 interclavicles, 2
cervical ribs
nMInG F21732 Steneosaurus edwardsi Cranium (anterior part),
lower jaw, 16 vertebrae (atlas-axis, 6 cervicals, 9 dorsals), 1
fifth metatarsal, calcaneum, rib fragments
nMInG F21731 Metriorhynchus superciliosus Upper and lower laws,
2 ilia, 28 vertebrae
nMInG F16892 Metriorhynchus superciliosus Skull
nMInG F21761 Peloneustes philacarpus Tooth (3 1)
nMInG F21762 Steneosauru sp. Tooth (3 1)
nMInG F21763 Steneosaurus sp. Tooth (3 1)
nMInG Unknown Tooth (3 2)—not located
Araujo.indd 29 14/01/2009 15:22:39
-
30 Irish Journal of Earth Sciences (2008)
paddle reconstructions, and never intended to send it as part of
the sale to Dublin. Indeed, it seems that the link between that
specimen and Dublin had been lost, insofar as the Leeds family was
concerned, by the time of Alfred Leeds’ death in 1917, the Eyebury
Register unusually records the number ‘77’ for Ophthalmosaurus as
reused without stating which institution the previous specimen went
to. Other paddle diagrams are included within the plates of E.T.
Leeds unpublished manuscript as figures, but this particular
diagram is unusual as it features the genus and number without
referring to the destination (either institution with accession
number, or Stürtz). Apart from a personal deal with Othniel Charles
Marsh in 1888, the Dublin sale was the first set of specimens
collected by the Leeds family that went to an institution other
than the BMnH in London, and it was to remain the only such example
until the commercial relationship was established with Stürtz in
1897. As such, it may simply be that Alfred Leeds was not yet in
the habit of recording which institution his specimens had gone to,
as everything up until then had gone to London.
A few questions remain regarding the Dublin Leeds Collection,
such as the exact horizon from which the material originated. The
numbers used by Leeds can often indicate a more specific location
or context from which the bones were collected through
cross-referring with the Eyebury Register, but unfortunately the
Dublin specimens are such an early part of what E.T. Leeds referred
to as the ‘Second Collection’ (Leeds 1956) it has only been
recorded that some of the Leeds numbers have been reused from
specimens sent to Dublin. This is not surprising, given that Alfred
Leeds states three years after the sale to Dublin that ‘I have no
memorandum of any of the bones … I am very sorry not to be able to
give you more information but I cannot remember what you had’
(Letter from Alfred Leeds, 28 October 1896—nMInH, October 1896
Correspondence File). This retrospective query from Dublin may well
have been the stimulus for Alfred Leeds to start keeping such
records for specimens that he subsequently collected, in the
Eyebury Register.
There are two types of labels applied by Leeds on Dublin
specimens: circular labels with large font numbers and oval labels
with small font numbers. Jeff Liston’s research in Glasgow had led
to the conclusion that the oval labels either represented a later
generation of the collection or the three figured (100 and above)
‘Miscellaneous’ specimens in the Eyebury Register—the Dublin
specimens, however, do not agree with these models. The majority of
the specimens have oval labels; only one specimen has solely round
labels, the Steneosaurus specimen nMInG F21732 (Leeds number
‘110’). not all the labelling is congruent,
however—confusingly the skull of specimen nMInG F21731
(Metriorhynchus) contains oval labels, whereas the postcranium and
lower jaw have round labels. More work on other Leeds specimens
will be necessary to determine if there are actually any real
patterns here, but Alfred Leeds’ 1896 letter states that he
‘think[s] the numbers were only put on to distinguish the sets’
(Letter from Alfred Leeds, 28 October 1896—nMInH, October 1896
Correspondence File), again indicating that the earlier
record-keeping practices of the collection had yet to evolve into
the rigorous form seen in the Eyebury Register.
The most taxonomically diverse group within the Dublin Leeds
Collection are the plesiosaurs—represented by three genera. This
agrees with typical Oxford Clay Formation reptile biodiversity
(Martill 1991).
Acknowledgements
Thanks to nigel Monaghan and Gareth Dyke for support and
discussion. Thanks also to Matthew Parkes and Graeme Lloyd for
assistance. Paolo Viscardi provided access to the stores and
Kimberly Eileen Lum interpreted handwritten correspondence. Mark
Evans and Mark Young gave additional comment on an early draft of
this paper. In particular, thanks are extended to Julian Leeds and
the Leeds family for access to and permission to use E.T. Leeds’
unpublished manuscript with the long-lost illustration of
‘Ophthalmosaurus 77’.
References
Adams-Tresman, S.M. 1987a The Callovian (Middle Jurassic) marine
crocodile Metriorhynchus from central England. Palaeontology 30,
179–94.
Adams-Tresman, S.M. 1987b The Callovian (Middle Jurassic)
teleosaurid marine crocodiles from central England. Palaeontology
30, 195–206.
Andrews, C.W. 1895 On the development of the shoulder girdle of
a plesiosaur (Cryptoclidus oxoniensis Phillips, sp.) from the
Oxford Clay. Annals and Magazine of Natural History 15, 333–46.
Andrews, C.W. 1910 A descriptive catalogue of the marine
reptiles of the Oxford Clay, part I. London. British Museum
(natural History).
Andrews, C.W. 1913 A descriptive catalogue of the marine
reptiles of the Oxford Clay, part II. London. British Museum
(natural History).
Arkell, W.J. 1933 The Jurassic system in Great Britain. Oxford.
Clarendon Press.
Baur, G. 1887 Ueber den Ursprungd er Extremitatend er
Ichthyopterygia. Berichte iiber die Versammlungend es
Oberrheinischeng eologischen Vereines 20, 17–20.
Benton, H.J. and Clark, J.M. 1988 Archosaur phylogeny and the
relationships of the Crocodylia. In M.J. Benton (ed.), The
phylogeny and classification of tetrapods, vol. 1, 295–338. Oxford.
Clarendon Press.
Araujo.indd 30 14/01/2009 15:22:41
-
Araújo et al.—The Alfred Leeds Collection of the NMINH 31
British Association for the Advancement of Science 1894 Report
of the Sixty-Third Meeting of the British Association for the
Advancement of Science, held at nottingham in September 1893.
London. John Murray.
Brown, D.S. 1981 The English Upper Jurassic Plesiosauroidea
(Reptilia) and a review of the phylogeny and classification of the
Plesiosauria. Bulletin of the British Museum (Natural History),
Geology 35, 253–347.
Caldwell, M.W. 1997 Limb osteology and ossification patterns in
Cryptoclidus (Reptilia: Plesiosauroidea) with a review of
sauropterygian limbs. Journal of Vertebrate Paleontology 17,
295–307.
Davis, J.W. 1888 Catalogue of the collection of fossils of
fossil fishes in the Science and Art Museum. Dublin. Dublin Museum
of Science and Art.
de Blainville, H.D. 1835 Description de quelques espèces de
reptiles de la Californie, précédée de l'analyse d'un system
general d'Erpetologie et d'Amphibiologie. Nouvelles Annales du
Muséum (National) d'Histoire Naturelle, Paris 4, 233–96.
de Blainville, H.D. 1853 Lettre de Monsieur de Blainville. In E.
Deslongchamps (ed.), Lettres sur les Crocodiles vivants et
fossiles, 103–38. Mémoires Société Linnéenne de normandie, 9.
Caen.
Duff, K.L. 1975 Paleoecology of a Bituminous shale—the Lower
Oxford Clay of central England. Palaeontology 18 (3), 443–82.
Eudes-Deslongchamps, J.A. 1868 Observations faites sur le foetus
du caïman à museau de brochet, et constatation d’un fait très
important dans l’anatomie des Crocodiles. Bulletin de la Societé
Linnéene de la Normandie 2(3), 222–5.
Eudes-Deslongchamps, J.A. 1867–9 Prodrôme des Téléosauriens du
Calvados. notes Paléontologiques. Caen 1, 9–354.
Fitzinger, L. 1843 Systema Reptilium. Vienna. Bramüller &
Seidel Ed.
Fraas, E. 1901 Die Meerkrkodile (Thalattosuchia n.g.) eine
Sauriergruppe der Juraformation. Jahreshefte des Vereins für
vaterländische Naturkunde in Württemberg 57, 409–18.
Gandola, R., Buffetaut, E., Monaghan, n., and Dyke, G. 2006 Salt
glands in the fossil crocodile Metriorhynchus. Journal of
Vertebrate Paleontology 26(4), 1009–10.
Geoffroy Saint-Hilaire, É. 1825 Recherches sur l’organisation
des Gavials, sur leurs affinités naturelles desquelles résulte la
nécessité d’une autre distribution générique. Gavialis,
Teleosaurus, Steneosaurus. Mémoires du Muséum Histoire Naturelle
12, 97–155.
Geoffroy Saint-Hilaire, É. 1831 Recherches sur de grands
sauriens trouvés à l´etat fossile aux confines maritimes de la
Basse-normandie, attribués d’abord au Crocodile, puis détermines
sous les noms de Teleosaurus et Steneosaurus. Mémoires de Academie
des Sciences 12, 1–138.
Gray, J.E. 1825 A synopsis of the genera of reptiles and
Amphibia, with a description of some new species. Annals of
Philosophy 26, 193–217.
Hay, O.P. 1930 Second bibliography and catalogue of the fossil
vertebrata of North America. Washington, DC. Carnegie Institute
Washington.
Kidston, R. 1888 Catalogue of the collection of Palaeozoic
plants in the Science and Art Museum. Dublin. Dublin Museum of
Science and Art.
Leeds, E.T. 1938/9 Eyebury and the Leeds Collection. Unpublished
MS., Leeds family papers.
Leeds, E.T. 1956 The Leeds Collection of fossil reptiles from
the Oxford Clay of Peterborough. Oxford. Blackwell.
Liston, J.J. and noè, L.F. 2004 The tail of the Jurassic fish
Leedsichthys problematicus (Osteichthyes: Actinopterygii) collected
by Alfred nicholson Leeds—an example of the importance of
historical records in palaeontology. Archives of Natural History
31, 236–52
Liston, J.J. and noè, L.F. 2008 ‘Old Bones’: The dinosaurs of
Alfred Leeds. In R. Moody, E. Buffetaut, D. Martill and D. naish
(eds), Dinosaurs (and other extinct saurians) a historical
perspective, 13–14. Abstracts of Meeting held on the 6-7 May 2008,
Geological Society London. London. Geological Society London.
Liston J.J. and Sanders H.L. 2005 The ‘other’ Glasgow boys: the
rise and fall of a school of palaeobotany. In A.J. Bowden, C.V.
Burek and R. Wilding (eds), History of palaeobotany: selected
essays, 197–228. London. Geological Society London.
Lydekker, R. 1884 Catalogue of vertebrate fossils from the
Siwaliks of India, in the Science and Art Museum, Dublin.
Scientific Transactions of the Royal Dublin Society 3, 69–86.
Lydekker, R. 1889 On the remains and affinities of five genera
of Mesozoic reptiles. Quarterly Journal of the Geological Society
45 (1–4), 41–59.
Lydekker, R. 1888–90 Catalogue of the fossil Reptilia and
Amphibia in the British Museum (Natural History), London. London.
British Museum (natural History).
Lydekker, R. 1891 Catalogue of the fossil mammal birds, reptiles
and amphibians in the Science and Art Museum. Dublin. Dublin Museum
of Science and Art.
Maisch, M.W. and Matzke, A.T. 2000 The Ichthyosauria.
Stuttgarter Beiträge zur Naturkunde 298B, 1–159.
Martill, D.M. 1991 Marine Reptiles. In D.M. Martill and J.D.
Hudson (eds), Fossils of the Oxford Clay. London. The
Palaeontological Association.
McGowan, C. and Motani, R. 2003 Ichthyopterygia. In Friedrich
Pfeil (ed.), Handbook of Paleoherpetology, part 8. Munich. Verlag
Dr. Friedrich Pfeil.
Monaghan, n.T. 1992 Geology in the national Museum of Ireland.
Geological Curator 5(7), 275–82.
neaverson, E. 1935 Reptiles from the Oxford Clay of Peterborough
in the geological collection of the University of Liverpool.
Proceedings of the Liverpool Geological Society 16, 234–62.
Owen, R. 1840 Report on British fossil reptiles. Report of the
British Association for the Advancement of Science 9, 43–126.
Phillips, J. 1871 The geology of Oxford and the valley of the
Thames. Oxford. Clarendon Press.
Sander, P. Martin 2000 Ichthyosauria: their diversity,
distribution, and phylogeny. Palaeontologische Zeitschrift 74,
1–35.
Seeley, H.G. 1869 Index to the fossil remains of Aves,
Ornithosauria and Reptilia, from the secondary system of strata
arranged in the Woodwardian Museum of the University of Cambridge.
Cambridge. Deighton, Bell & Co.
Seeley, H.G. 1874a Muraenosaurus leedsii, a plesiosaurian from
the Oxford Clay, part I. Quarterly Journal of the Geological
Society of London 30, 197–208.
Seeley, H.G. 1874b On the pectoral arch and forelimb of
Ophthalmosaurus, a new ichthyosaurian genus from the Oxford Clay.
Quarterly Journal of the Geological Society of London 30,
696–707.
Seeley, H.G. 1874c note on some of the generic modifications of
the Plesiosaurian pectoral arch. Quarterly Journal of the
Geological Society of London 30, 436–49.
Seeley, H.G. 1892 The nature of the shoulder girdle and
clavicular arch in Sauropterygia. Proceedings of the Royal Society
of London 51, 119–51.
Smellie, W.R. 1916 Apractocleidus teretipes: a new Oxfordian
plesiosaur in the Hunterian Museum, Glasgow University.
Transactions of the Royal Society of Edinburgh 51(19), 89–92.
Steel, R. 1973 Crocodylia, pt 16. In O. Kuhn (ed.), Encyclopedia
of palaeoherpetology. Stuttgart. Gustav Fischer Verlag.
Tarlo, B. 1960 A review of the Upper Jurassic pliosaur. British
Museum (natural History). Geology 4, 145–89.
Vignaud, P. 1995 Les Thalattosuchia, crocodiles marins du
Mésozoïque: systématique, phylogénétique, paléoécologie,
Araujo.indd 31 14/01/2009 15:22:43
-
32 Irish Journal of Earth Sciences (2008)
RICARDO ARAÚJODep. de Ciências da Terra, Fac. de Ciências e
Tecnologia, Univ. nova de Lisboa 2829–516 Caparica, Portugal.
ADAM STUART SMITH (corresponding author)Department of Geology,
Museum Building, Trinity College Dublin, Dublin 2, Ireland.
E-mail:[email protected]
JEFF LISTOnHunterian Museum,University of Glasgow, Glasgow,G12
8QQ,Scotland.
biochronologie et implications paléogéographiques. PhD thesis,
University of Poitiers, France.
von Meyer, H. 1830 Achte versammlung der naturforscher und
aerzte zu Heidelberg im September 1829. Mineralogische Abteilung,
15, 2 Isis v. Oken. 4/5/6, 517–19.
Walker, A.D. 1970 A revision of the Jurassic reptile Hallopus
victor (Marsh), with remarks on the classification of
crocodiles.
Philosophical Transactions of the Royal Society of London,
Series B, Biological Sciences 257 (816), 323–72.
Williston, S.W. 1925 The osteology of the reptiles. Cambridge,
MA. Harvard University Press.
Woodward, A.S. 1917 Obituary: Alfred nicholson Leeds, F.G.S.
Geological Magazine. Decade VI, 4, 478–80.
Araujo.indd 32 14/01/2009 15:22:44